Based on both phylogenetic hypothesis (BI and ML), H. vernayi is sister species to H. bayonii, presumably a relic in arid coastal ecosystems (Fig. 2, Fig. 6C), and forming a monophyletic group with H. bayonii, H. nzingae, and another species from northern Angola and Cabinda described below. The uncorrected p-distance found within H. vernayi for ND2 was 1.52%. The p-distances for ND2 between H. vernayi and their closely related species were: 14.15% with H. bayonii, 15.55% with H. nzingae, and 14.73% with the northern species described below (see Table 1). Small size and other morphological similarities have previously led to a misidentification of this taxon, or to its assignment to H. bayonii, by different authors.

ANGOLA • 2 ♀; Namibe Prov., Santa Marta, Lucira; –13.87861°, 12.42444°; 48 m a.s.l.; 24 May 2019; Pedro Vaz Pinto; FKH0226–7 and MNCN 50538 • 3 ♂; same collecting data as previous material; FKH0228, MNCN 50539, FKH0232 • 4 ♀, 1 ♂ same collecting data as previous material; 6 Jul. 2019; Pedro Vaz Pinto and Javier Lobón-Rovira; FKH0263–6, MNCN50533 • 1 ♀ juv.; same collecting information as previous material; FKH0268 • 2 ♀; Bentiaba; –14.17596°, 12.44689°; 150 m a.s.l.; 15 Feb. 2020; Pedro Vaz Pinto and Javier Lobón-Rovira; MNCN 50541 and ZMB 90448 • 2 ♂; same collecting information as previous material; FKH0417 and ZMB 90449 • 3 ♂; Lucira; September 1956; Charles Koch; TM 24447, 24450–1 • 1 ♂ and 1 ♀; Benguela Prov., Hanha; –12.24245°, 13.71399°; 39 m a.s.l.; 16 Nov. 2019; Pedro Vaz Pinto; ZMB 90450–1.

Hemidactylus vernayi is readily distinguished from non-Angolan congeners by sharing the same distinctive characters as H. bayonii (see Ceríaco et al. 2020a). In respect to the Angolan congeners, H. vernayi differs from H. longicephalus-group in its much smaller maximum size (maximum SVL 42.5 mm [mean=32.67] vs. 60.1 mm [mean=46.57] in H. longicephalus and 64.8 mm [mean=58.96] in H. paivae); from H. benguellensis by its lower number of precloacal pores (5–7 vs. 23–33); and from H. mabouia by having small subcaudal scales and a lower number of precloacal pores (5–7 vs. 28–39). Hemidactylus vernayi can further be distinguished from H. nzingae by fewer [11–16] rows of weakly keeled dorsal and caudal tubercles (vs. 16–21 rows of strongly keeled dorsal and caudal tubercles in H. nzingae). Interestingly, morphological analysis based on topotypic material of H. vernayi show extensive overlap with H. bayonii, and no clear diagnostic characters. Nevertheless, when we include the southern specimens of H. vernayi, these reveal a larger number of ventral scales across the belly (33–39 vs. 25–32 in H. bayonii, see Tables S3–4). See other new species descriptions for comparisons with this taxon.

In life (specimen FKH0417; Fig. 6A): dorsum dark brown with two lighter dorsolateral bands from the nares to the anterior part of the tail; the dorsal region presents a darker crossband from occipital to the medium the region of the tail where it fades before disappearing; the crossband in the dorsal region presents a W-shaped marking common in H. bayonii-group, which becomes V-shaped as it approach the posterior region; the head displays a linear crossband from the eye region to the occipital; all the crossbands along the body are separated by a light beige section; the ventral region has uniform beige coloration; supralabials have dark coloration while infralabials show the same light beige of the ventral region; limbs with irregular dark and light brownish patches; iris golden with a black narrow pupil and dark brown reticulation. Variation (Fig. 7): this species presents high variability in coloration from light whitish cream to brownish. Dorsal pattern can be present or absent. Furthermore, it was observed that in some individuals during the day the darker regions become lighter, making it often difficult to differentiate the dorsal pattern, due to a uniform light beige color across the whole body.

Figure 7. 

Dorsal photographs showing morphological variability found in Hemidactylus vernayi from (A) Hanha (ZMB 90450), (B) Bentiaba (MNCN 50541) and (C) Santa Marta (MNCN50533). Photos by Pedro Vaz Pinto (A) and Javier Lobón–Rovira (B, C).

This species is considered the sister species of H. bayonii, so far only known to occur along the arid coast from Bentiaba in northern Namibe Province, northwards to Hanha in coastal Benguela Province (Fig. 6). Despite being still poorly known, the species is probably widely distributed, and its habitat does not appear at present to be threatened; however, due to limited number of records we cannot calculate the extent of occurrence (EOO) and thus we regard the conservation status of the species as Data Deficient. Further studies are suggested to better assess its full distribution and conservation status.

Hemidactylus vernayi is an arid-adapted species, found foraging among small shrubs in arid coastal rocky semi-desert environments on the northern fringes of the Kaokoveld desert, and present also in vegetated valleys cutting through coastal semi-arid savannas. The species occurs along a coastal strip within the Semi-Arid Savanna AMBU, a region strongly influenced by the Namibe fog-belt (Cernak et al. 2012; Vaz Pinto et al. 2019). In the southernmost populations, most specimens were found associated with a recently described endemic small bush Commiphora benguellensis (Swanepoel 2015), while the northern populations were found in thicker and more diverse bush-dominated habitats. This species was always found foraging at night moving with agility between the spiny branches of small to medium-sized bushes, typically close to the ground and dropping quickly to the ground to seek shelter among crevices and cavities, or under small rocks or leaf-litter, when disturbed. Other species of geckos found in sympatry were Chondrodactylus fitzsimonsi, C. pulitzerae and Pachydactylus angolensis. No other Hemidactylus species were found occupying the same ecological niche as H. vernayi.

Based on both phylogenetic hypothesis (BI and ML), Hemidactylus pfindaensis sp. nov. is the sister group of a well-supported clade within the H. bayonii/nzingae-group, and represents a northern clade present in the Tropical and Subtropical Moist Forests Biome. Due to the lack of accurate morphological assessments within western African Hemidactylus until Ceríaco et al. (2020a), and the lack of detailed and extensive molecular data, this species may have been historically assigned to either H. longicephalus or H. muriceus, which have been reported from forest regions in northern Angola and Democratic Republic of Congo, respectively. However, morphological and genetic analyses, support H. pfindaensis sp. nov. as a new species (Table 2). The genetic ND2 p-distance differs 15.17% from H. bayonii, 17.21% from H. longicephalus and 14.01% from H. muriceus topotypic material.

ANGOLA • 1 ♀; Cabinda Prov., Chiloando; –5.12083°, 12.36667°; 95 m a.s.l.; 16 Mar. 2019; Pedro Vaz Pinto; FKH0178.

ANGOLA • 1 ♂; same collecting information as the holotype; without tail; MNCN 50537 • 1 ♂; Uíge Prov., Macocola; –7.01802°, 16.07658°; 952 m a.s.l.; 25 Sept. 2018; Pedro Vaz Pinto; without tail; FKH 0044 • 1 ♂; Uíge Prov., Serra do Pingano; –7.68451°, 14.92978°; 957 m a.s.l.; 31 Oct. 2014; Raffael Ernst; MTD 48932.

A medium sized Hemidactylus, with SVL of 45.53 mm (mean) with moderate long snout, 10 supralabials and 8–10 infralabials (Fig. 8). Dorsal pholidosis with 11–12 rows of moderate dorsal keeled tubercle scales and ventral pholidosis with 28–30 smooth scale rows on midbody. The species present a large, triangular mental scale, two large postmentals followed by two enlarged post-postmentals. Base of the tail with four large keeled dorsal tubercle rows and subcaudal scales small, about one fourth of the tail width. Males with 8 continuous precloacal pores. Five divided scansors beneath first digit of both manus and pes, seven beneath fourth digit of manus, seven or eight beneath the fourth digit of pes. Dorsum presents dark coloration with two light brown crossbands from the posterior part of the eye to the sacrum, where the two bands meet each other to form a V-shaped marking.

Figure 8. 

Holotype of Hemidactylus pfindaensis sp. nov. (FKH0178) from Chiloando, Cabinda Province, Angola. A – Dorsal and ventral view of body. B – Detail of head in lateral, dorsal and ventral views (from top to bottom). C – Detailed of left toes and right fingers (from left to right). Photos by Pedro Vaz Pinto.

Hemidactylus pfindaensis sp. nov. differs from the other non-Angolan, western and central African congeners, based on the same characteristics of the other Angolan species (Ceríaco et al. 2020a). However, this new species can easily be confused with H. muriceus-group, and differs from them by the presence of keeled dorsal tubercle scales vs. conical tubercle scales. Additionally, it differs from H. pseudomuriceus by lower number of precloacal-femoral pores (8 vs. 14–17) and one internasal scale vs. 2–3 in H. pseudomuriceus; and from H. echinus by the presence of considerably larger ventral than dorsal scales vs. similar size ventral and dorsal scales, and by reduced number of scansors underneath the 1^st and 4^th toe (5 and 7–8 vs. 10 and 12–13, respectively). Differentiated from H. steindachneri by lacking a longitudinal row of ventrolateral keeled tubercles, and from H. hecqui in not having the nostrils in contact with the first supralabial. Hemidactylus pfindaensis sp. nov. can be distinguished from H. mabouia by the presence of smaller subcaudal scales (large and elongated in H. mabouia) and from H. benguellensis by lower number of precloacal-femoral pores (8 vs. 23–33), and the dark dorsal color with dorsolateral light stripes and absence of dorsolateral orange tubercle rows (present in H. benguellensis). Hemidactylus pfindaensis sp. nov. differs from H. longicephalus-group by having a smaller SVL (maximum length 49.09 mm [mean=45.53] vs. 60.08 [mean=46.57] in H. longicephalus and 64.8 [mean=58.96] in H. paivae), more keeled tubercle scales and a lower number of dorsal tubercle rows (12 vs. 13–17 in H. longicephalus, and 13–16 in H. paivae). It differs from the H. bayonii-group by having larger SVL (maximum length 49.09 mm [mean=45.53]), than H. bayonii 36.2 mm [mean=34.9] and H. vernayi (42.5 mm [mean=32.89]); from H. bayoni by having lower number of dorsal tubercle rows (12 vs. 14–16 in H. bayonii), and head more compressed (HL/HW [mean=1.7] in H. bayonii vs. HL/HW [mean=1.5] in H. pfindaensis sp. nov.); from H. vernayi by presence of more precloacal-femoral pores (8 in H. pfindaensis sp. nov. vs. 4–6 in H. vernayi), lower number of ventral scales across the belly (28–30) than H. vernayi (32–39) and higher number of infralabials (9–10 vs. 7–8); and differs from H. nzingae and H. gramineus by larger number of scales across the belly (28–30 vs. 22–27 and 23–25, respectively), lower number of dorsal tubercle rows than H. nzingae (12 vs. 16–21) and larger number of granular scales between the dorsal tubercle rows than H. nzingae and H. gramineus (4–5 vs. 2–3).

Measurements and meristic characters of the holotype are presented in Table S5. Adult female with a snout-vent-length (SVL) of 49.09 mm, a regenerated tail length (TL) of 32.93 mm. Body slender, nape distinct. Head slightly narrower than the body and largely elongated (HW/HL 0.62). Canthus rostralis not prominent, but well-marked. Eye diameter (2.96 mm), with vertical pupil and crenulated margin. Supraciliar scales small and slightly pointed. Ear height (0.93 mm). Ear to eye distance slightly larger than orbit diameter (3.6 mm). Snout rounded. Frontal scales granular and larger than occipital scales. Occipital scales granular with lateral conical a large tubercle scale. Rostral wider than deep (2.21 vs. 1.04 mm, respectively). Rostral semidivided anterodorsally, in contact with 1^st supralabial, nostril, prenasal and one internasal scales. 11 supralabials and 11 infralabials. First supralabial in contact with the nostril. Nostril circular rounded by rostral, supranasal, two postnasal and first supralabial. Postnasals larger than supranasal. Nostril in direct contact with the rostral and 1^st supralabial. One row of scales between supralabials and the orbit. Mental large and markedly triangular, with two large rectangular postmental scales in broad contact posteriorly to the mental. 5 post-postmental scales, composed by post-postmental slightly smaller than postmental scales in contact with postmentals, 1^st and 2^nd infralabials, and 3 small post-postmental in contact with postmental scales. Gular scales and granular smaller than ventral scales. Between the gular scales and infralabials, a row of enlarged scales is present, decreasing in size until the 5^th infralabial where they become the same size as the gular scales.

Body relatively slender and elongated (TRL/SVL 0.45). Ventral scales about double size than dorsal scales, with 28 scales across the belly. The dorsal pholidosis present heterogenous conical, granular scales interspersed by 12 keeled dorsal tubercle rows at midbody. Dorsal tubercle rows are separated by 4–5 granular scales. Tubercle scales reach the posterior part of head and the nape, where tubercle scales lose the keeling progressively. Base of the tail with four large keeled dorsal tubercle rows dorsally and subcaudal scales small, about one fourth of the tail width. Regenerated tail has no presence of tubercle scales, having largely homogeneous scales along and across the tail. Precloacal scales enlarged and one well-developed postcloacal spurs on each side.

Fore- and hindlimbs relatively short, stout; forearm short (FL/SVL 0.17); tibia short (CL/SVL 0.18). Short digits and clawed. All digits of manus and pes indistinctly webbed. Scansors beneath each toe equally divided, with the exception of 1^st and terminal scansor undivided. 4^th and 5^th toes with 2 and 3 undivided terminal scansors, respectively. Scansors beneath each finger equally divided, with the exception of 1^st and two terminal scansor undivided. 1^st and 5^th fingers with 3 undivided terminal scansors. Number of scansors: 5-7-7-8-7 (right manus), 5-7-8-8-9 (right pes). Relative length of digits: V < IV=III=II > I (right manus); V < IV=III > II > I (right pes).

Variation in scalation and body measurements of the paratypes of H. pfindaensis sp. nov. are reported in Table S5. All the material analyzed agrees entirely with the holotype description. However, paratype FKH0179, shows undivided rostral scales and MTD 48932, 3^rd and 4^th supralabials semi-fused at the base.

In life (specimen FKH0178; Fig. 9A): this species presents dark coloration over the dorsum with two light brown crossbands from the posterior part of the eye to the sacrum, where the two bands meet each other to form a V-shaped marking; the rest of the dorsal part of the body is mostly dark uniformly brownish with some patches of light brown, especially at the nostril area and the hindlimbs. Coloration of the regenerated tail is uniform dark brown. The ventral part of the body is lighter, fully covered with scattered black speckles, from head to tail. Iris golden with a black narrow pupil and brownish-golden reticulation. In preservative (Holotype; Fig. 8): dorsum with dark uniform coloration; ventrum is light beige with scattered black speckles. Variation: occasionally difficult to differentiate the dorsal pattern, due to a uniform dark coloration across the whole body.

Figure 9. 

A – Dorsal view of Hemidactylus pfindaensis sp. nov. (Holotype, FKH 0178). B – habitat of H. pfindaensis sp. nov. C – records of H. pfindaensis sp. nov. (yellow circles represent records of H. pfindaensis sp. nov.; star depicts the type locality). Photos by Pedro Vaz Pinto (A) and Luke Verburgt (B).

The name “pfindaensis” derives from the local word “pfinda” which in Kikongo – the main language used in Uíge Province and northwestern Angola – refers to a “gallery forest” or a “continuous block of thick forest”, the main habitat type associated with the species.

A typical forest gecko, this species has been found in two sites of northern Angola, both in Uíge Province, and on a third site in the enclave of Cabinda. Its known presence north and south of the Congo River, suggests a much larger distribution range, that will likely extend to Democratic Republic of the Congo and Republic of the Congo, and possibly also into Gabon. In Angola, it will also likely be present in the Mayombe Forest and Northeastern Forest-Savanna Mosaic. However, due to limited number of records we cannot calculate the EOO and thus we regard the conservation status of the species as Data Deficient, and further studies are suggested to better assess its full distribution and conservation status. Although central African forests are currently threatened by deforestation and human encroachment, the occurrence of this species across a large geographical and altitudinal range, suggests that it is likely common, yet further studies are necessary to evaluate its conservation status.

Hemidactylus pfindaensis sp. nov. appears to be a species strongly associated with moist evergreen forests. Specimens were collected at various altitudes, both near sea level and above 900 m a.s.l., but always in moist gallery forest, within the Northwestern Forest-Savanna Mosaic. All specimens were found foraging at night on tree trunks of well-developed trees, approximately 1–2 m above ground. Although never found hiding, it seems likely that it finds shelter under tree bark.

The original description of H. benguellensis by Bocage make reference to “… species with 16 to 18 longitudinal series, 9 upper labials and 9 lower and 26 pre-cloacal and femoral pores, 13 on each side in a continuous series”. These characters were later corroborated by Monard (1937), with one male from Ebanga differing only from Bocage material on the number of preclocal-femoral pores (28 vs. 26).

In February 2020, a survey was conducted at Ebanga, Benguela Province, about 40 kilometers south of Cahata, the original type locality (Bocage 1893), and from where some of the earliest material ascribed to this species had been collected (Monard 1937). During this expedition an adult male morphologically assigned to H. benguellensis was collected on a granite rock boulder. A few days after, a second male was collected on a rock face overhanging a stream, near Passe, Benguela Province, ~80 km west from Ebanga and the type locality (Table S2, Fig. 12).

Figure 12. 

A – From top, dorsolateral view of Hemidactylus cinganji sp. nov. (FKH0435), H. benguellensis (FKH0413) and H. carivoensis sp. nov. (MNCN 50543); B – records of H. benguellensis–group (purple color represents records of H. cinganji sp. nov.; blue H. benguellensis; dark green H. carivoensis sp. nov.; circles represent genetically assigned records, while triangles represent historical or non-genetically assigned records by Ceríaco et al. 2020a; color stars depict type or neotype localities; black star depicts original type locality of H. benguellensis assigned by Bocage (1893); C – habitat of H. cinganji sp. nov. at Ebanga; D – habitat of H. carivoensis sp. nov between Dombe–Echimina. E – habitat of H. benguellensis at Tchivinguiro. Photos Javier Lobón–Rovira (A–D) and William R. Branch (E).

Interestingly, our phylogenetic analysis revealed a genetically separate lineage well-differentiated from the material assigned to H. benguellensis by Ceríaco et al. (2020a) (Fig. 2), with uncorrected p-distance 11.64% (Table 1). However, the available historical material collected between 1925 and 1953, from Ebanga, Hanha and Entre Rios (Ceríaco et al. 2020a), was morphologically included as part of the taxon genetically assigned to H. benguellensis by Ceríaco et al. (2020a), without any molecular support. It should be noted that, not only it is unknown to which phylogenetic clade should the material from Hanha and Entre Rios be assigned to, but the problem is further compounded by some geographic inconsistencies. The specimens from Hanha, obtained during the VAE were most likely collected at Hanha Estate (also known as Hanha do Norte) and located on the semi-arid coast north of Lobito (see Branch et al. 2017 for a detailed discussion on misinterpretations surrounding the toponym Hanha), and not on the geographic location given by Ceríaco et al. (2020a), about 130 km to the southeast and on the base of the escarpment at 917 m a.s.l. On the other hand, the locality of Entre Rios (meaning “between rivers”) can also be confusing, as several historical toponyms in Angola bear that name, and various publications have placed Entre Rios in different locations, ranging from 1,200 to 1,500 m a.s.l. in Benguela and Huambo provinces (e.g. Conradie et al. 2013; Marques et al. 2018; Ceríaco et al. 2020a). On his first Angolan expedition, Hellmich established his main base at the farm of Entre Rios, owned by a German citizen named Alfons Burger (Hellmich 1957), which is located further south than previously acknowledged, at –13.30385° 14.70761°, 1076 m a.s.l, Benguela Province. Two specimens were here collected by Hellmich in 1953 and a third specimen was added by Herr Burger in the following year, all initially identified as H. mabouia, and only recently reassigned to H. benguellensis (Ceríaco et al. 2020a). Nevertheless, the lack of phylogenetic context, the morphological variability recognized for specimens belonging to the neotypic clade, small sample size of the topotypic material, and the fact that Hanha and Entre Rios are intermediately located in relation to specimens genetically ascribed to different clades, recommends caution before assigning those specimens.

Based on field observations, the material assigned to H. benguellensis by Ceríaco et al. (2020a) is a very variable species present in different biomes and biogeographic units and occupying different niches, often showing arboreal habits, frequently found on the trunk or branches of trees, but sometimes in rocks, and also around human infrastructures such as man-made walls. In contrast, the new linage revealed here from the topotypic region, was only found associated with large granitic boulders in mountain habitats, even though it is a very limited sample.

Furthermore, the material recently assigned to H. benguellensis by Ceríaco et al. (2020a) presents some incongruence with our new topotypic material and the original H. benguellensis description. The historical material collected from Cahata and Ebanga (Bocage 1893; Monard 1937), makes reference to large sized specimens (SVL 55 mm and 42 mm, respectively), with 9 supralabials and 26–28 precloacal-femoral pores. Remarkably, this description agrees fully with the new specimens collected in Ebanga and Passe while underscoring differences from the recently reassigned H. benguellensis, which consistently presents 10–11 supralabials (Table S6). Moreover, the neotype specimen that was ascribed to H. benguellensis by Ceríaco et al. (2020a) is an adult male with 40.8 SVL and 33 precloacal-femoral pores. Therefore, based on a morphological evidence and incongruence with the original description, clear well-differentiated genetic distance and morphological differences between the two clades were detected, adding to the large distance from the original type locality (>250 km). Notably, the new specimens found near the original topotypic agrees perfectly with the original H. benguellensis described by Bocage (1893) and later reported by Monrad (1937) from Ebanga. However, due to the recent redescription and new type locality assigned by Ceríaco et al. (2020a), and in order to maintain taxonomic stability and to avoid more confusion within this group, we consider the new material recovered from near the original type locality of H. benguellensis as a new species.

ANGOLA • 1 ♂; Benguela Prov., Ebanga; –12.77082°, 14.77102°; 1916 m a.s.l; 22 Feb. 2020; Pedro Vaz Pinto and Javier Lobón-Rovira; FKH0439.

ANGOLA • 1 ♂; Benguela Prov., Passe; –12.66774°, 14.01607°; 828 m a.s.l; 13 Feb. 2020; Pedro Vaz Pinto and Javier Lobón-Rovira; FKH0413.

A large sized Hemidactylus, with maximum SVL of 55 mm (mean) and maximum width of 10.5 mm (Fig. 13). Nine supralabials and 9–11 infralabials. Dorsal pholidosis with 13–18 rows of moderate strongly keeled tubercle scales and ventral pholidosis with 33–41 smooth and rounded scale rows on midbody. Hemidactylus cinganji sp. nov. present a moderate, triangular mental scale, two large postmentals followed by two medium post-postmentals. Tail with six strongly keeled dorsal tubercles rows dorsally and subcaudal scales medium sized, interspersed by 11 horizontal whorls of keeled scales on the original portion of the tail. Males with 26–28 continuous precloacal-femoral pores and 1–4 postcloacal tail spurs. Six divided scansors beneath first digit of both manus and pes, seven or eight beneath fourth digit of manus, eight or nine beneath the fourth digit of pes. Dorsum presents dark orange-beige coloration across the body with darker coloration on the dorsal part of the head that continues along the medial part of the dorsum until the sacrum.

Figure 13. 

Holotype of Hemidactylus cinganji sp. nov. (FKH0439) from Ebanga, Benguela Province, Angola. A – Ventral and dorsal view of body. B – Detailed of head in lateral, dorsal and ventral views (from top to bottom). Photos by Pedro Vaz Pinto (A) and Rogério Ferreira (B).

Hemidactylus cinganji sp. nov. can be distinguished from other non-Angolan western and central Africa congeners based on the same characteristics of H. benguellensis (Ceríaco et al. 2020a). Hemidactylus cinganji sp. nov. can be distinguished from H. mabouia by the presence of smaller subcaudal scales (vs. large, elongated scales). It differs from the H. bayonii-group by its large size, largely keeled dorsal tubercle scales vs slightly keeled and larger number of precloacal-femoral pores (26–28 vs. 4–9 in H. bayonii, 4–6 in H. vernayi, 7–8 in H. nzingae and 8 in H. gramineus). It differs from H. longicephalus-group by larger number of precloacal-femoral pores (26–28 vs. 6–11 in H. longicephalus and 6–8 in H. paivae). It differs from H. benguellensis by lower number of supralabials (9 vs. 10–11 in H. benguellensis).

Measurements and meristic characters of the holotype are presented in Table S6. Adult male with a snout-vent-length (SVL) of 45.05 mm and a tail length (TL) slightly smaller than SVL 42.07 mm. Body slender, nape distinct. Head slightly narrower than the body and moderate head length (HW/HL 0.72). Canthus rostralis not prominent. Eye diameter (3.05 mm), with vertical pupil and crenulated margin. Supraciliar scales small and rounded. Ear height (1.47 mm). Ear to eye distance slightly larger than orbit diameter (4.17 mm). Snout rounded and pointed. Frontal scales granular and larger than occipital scales. Occipital scales granular interspersed with large number of smooth and keeled tubercle scales from eyes to nape. Rostral undivided, in contact with 1^st supralabial, nostril, supranasal and one internasal scales. 9 supralabial and 8–9 infralabials. First supralabial and rostral in direct contact with the nostril. Nostril circular rounded by rostral, 1^st supralabial, supranasal, and two postnasals. One row of scales between supralabials and the orbit. Mental large, triangular and rounded posteriorly, with two larges rectangular postmental scales in broad contact posteriorly to the mental. 2 post-postmental scales, in contact with post-postmental slightly smaller than postmental scales and 1^st and 2^nd infralabials. Gular scales slightly smaller than ventral scales and granular.

Body relatively robust and slightly elongated (TRL/SVL 0.40). Ventral scales widely larger than dorsal scales, with 32 scales across the belly. The dorsal pholidosis present heterogenous conical, granular scales interspersed by 15 strongly keeled dorsal tubercle rows of at midbody. Dorsal tubercle rows are separated by 3 granular scales. Tubercle scales reach the posterior part of the eye region where tubercle scales remain keeled. Tail with six strongly keeled dorsal tubercles rows dorsally and subcaudal scales medium sized, interspersed by 11 horizontal whorls of keeled scales on the original portion of the tail. Regenerated portion of the tail presents homogeneous scales all around the surface without tubercle scales. 27 precloacal-femoral pores enlarged and 2 well-developed postcloacal spurs on each side.

Fore- and hindlimbs relatively short, stout; forearm short (FL/SVL 0.15); tibia short (CL/SVL 0.16). Short digits strongly clawed. All digits of manus and pes indistinctly webbed. Scansors beneath each toe composed by 1^st and terminal scansor undivided with the exception of 4^th and 5^th toes with 3 and 2 undivided terminal scansor, respectively. Scansors beneath each finger equally divided, with exception of 1^st and last terminal scansor undivided. Number of scansors: 5-5-6-7-6 (right manus), 6-8-8-9-7 (right pes). Relative length of digits: V < IV=III > II > I (right manus); V < IV < III > II > I (right pes).

Variation in scalation and body measurements of the holotype and paratype of H. cinganji sp. nov. are reported in Table S6.

In life (holotype FKH0439; Fig. 12A): this species displays a dark orange-beige coloration across the body with darker coloration on the dorsal part of the head that continues along the medial part of the dorsum until the sacrum; in the dorsal section it presents two lighter cream dorsolateral bands from the nares to the tail; the ventral part of the head shows a cream coloration with dispersed black speckles that increase in the anterior part of the infralabials; ventrum is uniform light beige or cream; supralabials are darker than infralabials; fore- and hindlimbs with irregular dark brownish coloration; tail with similar color and slightly banded; iris silvered with a black narrow pupil and brownish-golden reticulation. In preservative (Holotype; Fig. 13): dorsum with dark brownish coloration and two lighter dorsolateral bands from the narine till the sacrum; ventrum is light uniform beige. Variation: no variation was observed.

The name “cinganji” is a widespread traditional word used in Angolan local languages that represents an ancestral spiritual entity that reincarnates assuming different physical forms in different places and occasions. This name is suitable as the new species corresponds to a taxon that was first described under a different name, then became lost and now resurfaces after its original name had been hijacked by a surrogate sister-species. The species epithet is used as a neuter singular noun in opposition to the generic name.

The full distribution range of the species remains unknown, even though so far it has only been confirmed to occur in the interior of Benguela Province, from Cahata to Passe and Ebanga, above 800 m a.s.l. Due to poor sampling and few material confirmed to belong to this species, we cannot calculate the EOO and thus we regard the conservation status of the species as Data Deficient. Additional data will need to be collected to address its conservation status.

Hemidactylus cinganji sp. nov exhibited a rock-dwelling behavior associated to large granitic boulders in mountainous regions from medium to high altitudes. The confirmed records fall in Benguela’s transition region between the Scarp and Transitional Zone and the Angolan Highlands. Both specimens were collected at night while foraging in large vertical granitic walls, one on a mountain cliff and the other on a boulder overhanging a forest stream. However, due to the poor sampling of this species, its natural history and real habitat requirements remain uncertain.

Hemidactylus carivoensis sp. nov. represents a new form genetically divergent from the two related species, with ND2 p-distance of 12.51% from H. cinganji sp. nov. and 12.48% from H. benguellensis (Table 1). Given the hypervariable morphology of the H. benguellensis-group, and the lack of molecular and morphological information before Ceríaco et al. (2020a), it is possible that some historical records of H. benguellensis may be assignable to this species. However, the morphological characteristics of H. carivoensis sp. nov. allows it to be unequivocally distinguished from H. benguellensis and H. cinganji sp. nov. (Table 2). Moreover, H. benguellensis has not yet been reported within the known distribution range of this newly described species (Marques et al. 2018; Ceríaco et al. 2020a; this study).

ANGOLA • 1 ♀; Benguela Prov., Fazenda Carivo; –13.19167°, 13.41806°; 382 m a.s.l.; 15 Jul. 2018; Pedro Vaz Pinto; good condition with a ventral-lateral incision for the removal of liver sample and some torn skin in the ventrum; FKH0033.

ANGOLA • 1 ♀, juv.; Benguela Prov., Santa Maria, Praia do Meva; –13.19167°, 12.99139°; 287 m a.s.l.; 12 Nov. 2016; Pedro Vaz Pinto and William R. Branch; PEM R24218 (field number AG 16.63) • 1 ♂; same collecting locality as holotype; 7 Jan. 2020; MNCN 50542 • ♂; same collecting locality as holotype; Dec. 2020; ZMB 90453 • 1 ♂; Benguela Prov., between Dombe and Equimina; –13.18333°, 12.99139°; 367 m a.s.l.; 14 Feb. 2020; Pedro Vaz Pinto and Javier Lobón-Rovira; MNCN 50543 • 1 ♀; same collecting details as previous material; ZMB 90452.

ANGOLA • 2 ♂, subadult; same collecting locality as holotype; Dec. 2020; MNCN 50544 and FKH0499 • 1 ♀; same collecting locality as previous; FKH0500.

A medium sized Hemidactylus, with SVL of 43.2 mm (mean) and maximum width of 8.8 mm (Fig. 14). 9–11 supralabials and 8–10 infralabials. Dorsal pholidosis with 15–22 rows of moderate strongly keeled tubercle scales and ventral pholidosis with 32–38 smooth and rounded scale rows around midbody. Hemidactylus carivoensis sp. nov. present a moderate, triangular, moderate rounded posteriorly mental scale, two large postmentals followed by two large post-postmentals. Tail with four strongly keeled dorsal tubercles rows dorsally and subcaudal scales small, about one fourth of the tail width, interspersed by large series of horizontal whorls of keeled scales. Males with 15–22 continuous precloacal-femoral pores. Five divided scansors beneath first digit of both manus and pes, seven or eight beneath fourth digit of manus, nine beneath the fourth digit of pes. Dorsum darker coloration on the dorsal part of the head that continues along the medial part of the dorsum until the sacrum.

Figure 14. 

Holotype of Hemidactylus carivoensis sp. nov. (FKH0033) from Carivo, Benguela Province, Angola. A – Ventral and dorsal view of body. B – Details of head in lateral, dorsal and ventral views (from top to bottom). C – Details of right toes and left fingers (from left to right). Photos by Pedro Vaz Pinto.

Hemidactylus carivoensis sp. nov. can be distinguished from other non-Angolan western and central African congeners based on the same characteristics of H. benguellensis (Ceríaco et al. 2020a). Hemidactylus carivoensis sp. nov. can be distinguished from H. mabouia by the presence of smaller subcaudal scales (vs. large elongated scales). It differs from the H. bayonii-group by the presence of largely keeled dorsal tubercle scales vs. slightly keeled, larger number of precloacal-femoral pores (17–22 vs. 4–9 in H. bayonii, 4–6 in H. vernayi, 7–8 in H. nzingae and 8 in H. gramineus), larger number of ventral scales across the belly (34–38) than H. bayonii (29–32), H. nzingae (22–27), H. gramineus (23–25) and H. faustus sp. nov. (29–32). The new species differs from H. longicephalus-group by a larger number of precloacal-femoral pores (17–22 vs. 6–11 in H. longicephalus and 6–8 in H. paivae), smaller SVL (maximum SVL 45.2 mm [mean=43.19] in H. carivoensis sp. nov. vs. 60.07 mm [mean=46.6] in H. longicephalus and 68.4 mm [mean=58.96] in H. paivae) and lower number of lamellae under 1^st toe (5 in H. carivoensis sp. nov. vs. 6–7 in H. longicephalus-group). Hemidactylus carivoensis sp. nov. differs from H. benguellensis-group by less keeled and rounded dorsal tubercles, number of precloacal-femoral pores (15–22 vs. 23–33 in H. benguellensis and 26–28 in H. cinganji sp. nov.), the presence of consecutive series of keeled subcaudal scales under the tail whorls, larger number of ventral scales across the belly (34–38 in H. carivoensis sp. nov. vs. 30–32 in H. cinganji sp. nov.) and large number of supralabials (10–11 in H. carivoensis sp. nov. vs. 9 in H. cinganji sp. nov.) (see Table 2, for comparison with other Angolan congeners).

Measurements and meristic characters of the holotype are presented in Table S7. Adult female with a snout-vent-length (SVL) of 45.16 mm and a tail length (TL) about the same size than SVL 44.79 mm. Body slender, nape distinct. Head slightly narrower than the body and largely elongated (HW/HL 0.65). Canthus rostralis not prominent. Eye diameter (3.04 mm), with vertical pupil and crenulated margin. Supraciliar scales small and rounded. Ear height (0.83 mm). Ear to eye distance slightly larger than orbit diameter (3.46 mm). Snout rounded and pointed. Frontal scales granular and larger than occipital scales. Occipital scales granular interspersed with large number of smooth and conical tubercle scales from eyes to nape. Rostral wider than deep (2.31 vs. 0.93 mm, respectively). Rostral semi-divided posterodorsally, in contact with 1^st supralabial, nostril, two postnasals and one internasal scales. Eleven supralabials and 9 infralabials. First supralabial in contact with the nostril. Nostril circular rounded by rostral, supranasal and two postnasals. Postnasals larger than supranasal. One or two rows of scales between supralabials and the orbit. Mental large, triangular and rounded posteriorly, with two large rectangular postmental scales in broad contact posteriorly to the mental. 4 post-postmental scales, composed by post-postmental slightly smaller than postmental scales in contact with postmentals and 1^st and 2^nd infralabials, and 2 post-postmental half size than lateral post-postmental in contact with postmental scales. Gular scales slightly smaller than ventral scales and granular. Between the gular scales and infralabials a row of enlarged scales is present, decreasing in size until the 5^th infralabial where they become the same size as the gular scales.

Body relatively robust and slightly elongated (TRL/SVL 0.39). Ventral scales widely larger than dorsal scales, with 36 scales across the belly. The dorsal pholidosis present heterogenous conical, granular scales interspersed by 17 strongly keeled dorsal tubercle rows of at midbody. Dorsal tubercle rows are separated by 3 granular scales. Tubercle scales reach the posterior part of the eye region where tubercle scales lose the keeling progressively. Tail with four strongly keeled dorsal tubercles rows dorsally and subcaudal scales small, about one fourth of the tail width, interspersed by 22 horizontal whorls of keeled scales. Precloacal scales enlarged and one well-developed postcloacal spur on each side.

Fore- and hindlimbs relatively short, stout; forearm short (FL/SVL 0.16); tibia short (CL/SVL 0.18). Short digits and clawed. All digits of manus and pes indistinctly webbed. Scansors beneath each toe composed by 1^st and two terminal scansor undivided. 4^th toes with 3 undivided terminal scansors. Scansors beneath each finger equally divided, with the exception of 1^st and last terminal scansors undivided. Number of scansors: 6-6-7-7-7 (right manus), 7-8-9-10-7 (right pes). Relative length of digits: V < IV=III > II > I (right manus); V < IV < III > II > I (right pes).

Variation in scalation and body measurements of the paratypes of H. carivoensis sp. nov. are reported in Table S7. All the material analyzed is in agreement entirely with the holotype.

In life (specimen MNCN 50543; Fig. 12A): this species displays an orange-beige coloration across the body with darker coloration on the dorsal part of the head that continues along the medial part of the dorsum until the sacrum; in the dorsal section presents two lighter dorsolateral bands from the nares to the tail; the ventral part of the head shows a cream coloration with dispersed black speckles that increase towards the anterior part of the infralabials; ventrum is uniform light beige or cream; supralabials are darker than infralabials; fore- and hindlimbs present an irregular dark brownish coloration; tail with similar color and slightly banded; iris silvered with a black narrow pupil and brownish-golden reticulation. In preservative (Holotype; Fig. 14): dorsum with dark greyish coloration and two lighter dorsolateral bands from the narine till the sacrum; ventrum is light uniform beige. Variation: ventrum can be uniform beige or present scattered black speckles; crossband of the tail can be present or absent.

The species epithet “carivoensis” refers to the Farm Carivo, an old estate situated along the banks of the mid-lower Coporolo River on the coastal plain of Benguela Province, and where most of the type series was collected. The species proved to be common in the area, and by recognizing the farm, we also acknowledge the ongoing support from the owners to researchers, similar to the Chapmans nearly a century ago.

The full distribution range of the species remains unknown, even though so far it has only been confirmed to occur across the coastal plain of western Benguela Province. Due to the variable morphology of the taxon known as H. benguellensis and which formerly included material now assigned to H. cinganji sp. nov., it is possible that some historical material was wrongly identified and should instead belong to this newly described species. Spiny savannas are well distributed in western Angola, suggesting that the new species might be common across a wider range than what is currently known, but available data is still poor. However, due to limited material confirmed to belong to this species, we can’t calculate the EOO and we regard the conservation status of this species as Data Deficient. Therefore, it is important for future studies to establish the real distribution of H. benguellensis, H. cinganji sp. nov. and H. carivoensis sp. nov. to better determine the conservation status of the species.

Hemidactylus carivoensis sp. nov. represents an arboreal species occurring on coastal savanna habitats dominated by small acacia (Senegalia spp.) trees and bushes, and often with abundant accumulation of dead wood. Their preferred habitat is well contained within the Semi-arid Savannas. The species was found during night surveys and in sympatry with Afrogecko ansorgii, albeit in different niches. While individuals of A. ansorgii were found foraging in small twigs and branches at medium height of bushes (Vaz Pinto et al. 2019), H. carivoensis sp. nov. were always collected at the base of bushes and tree trunks, or in fallen branches or dead wood on the ground.

The phylogenetic analysis revealed that Hemidactylus faustus sp. nov. clusters within a large clade, which includes the H. nzingae-group, H. bayonii-group, H. benguellensis-group, and H. pfindaensis, although its phylogenetic position is not well-stablished (Fig. 2). However, according to the morphological and genetic analysis performed, this species represents a well-differentiated clade from H. bayonii-group and H. benguellensis-group, with 17.85% and 16.72% uncorrected p-distance, respectively (Table 1). This species represents a micro-endemic form only known from a unique geological formation, possibly a relic species confined to the massive conglomerate inselbergs of Pungo Andongo and surrounds, in Malanje Province. Due to their exclusive morphological characteristics and the elusive behavior of the species, we consider that no other specimens have been reported before and mistaken with any of its congeners.

ANGOLA • 1 ♀; Malanje Prov., Pungo Andongo; -9.67333°, 15.59222°; 1217 m a.s.l.; 11 Jul. 2019; Pedro Vaz Pinto and Javier Lobón-Rovira; good condition with partially regenerated tail; FKH0281.

ANGOLA • 1 ♂; same collecting details as holotype; MNCN50534 • 2 ♀; same collecting details as holotype; MNCN50535 and ZMB 90447 • 1 ♀; same collecting locality as holotype; 11 Aug. 2018; Beatriz Vaz Pinto; FKH0023.

ANGOLA • 1 ♀; same locality as type material; 11 Aug. 2018; Pedro and Afonso Vaz Pinto; ZMB 90446 • 1 ♂, juv.; same collecting details as previous material; ZMB 90445 • ♀; same locality as type material; 15 Oct. 2020; Pedro Vaz Pinto; MNCN 50536.

A robust medium sized Hemidactylus, with SVL of 39.4 mm (mean) and maximum width of 7.4 mm (Fig. 15). 8–9 supralabials and 7–8 infralabials. Dorsal pholidosis with 15–17 rows of moderate keeled tubercle scales and ventral pholidosis with 29–32 smooth and rounded scale rows around midbody. Hemidactylus faustus sp. nov. present a moderate, triangular mental scale, two large postmentals followed by two large post-postmentals. Tail with thickness at the base tail with conical tubercle rows laterally. Regenerated tail with regular larger scales. Males with 17–19 continuous precloacal-femoral pores. Five or six divided scansors beneath the first digit of both manus and pes, seven beneath the fourth digit of manus, eight or nine beneath the fourth digit of pes. Dorsum coloration with two darker dorsolateral bands from the occiput to the tail, which includes 5 W-shaped darker crossbands in contact with both lateral bands; each dark crossbar is separated by lighter blotches.

Figure 15. 

Holotype of Hemidactylus faustus sp. nov. (FKH0281) from Pungo Andongo, Malanje Province, Angola. A – Ventral and dorsal view of body. B – Details of head in lateral, dorsal and ventral views (from top to bottom). C – Details of left toes and right fingers (from left to right). Photos by Pedro Vaz Pinto.

Head slightly more quadrangular than the other members of the Angolan Hemidactylus (HL/HW ≤1.4 vs. >1.5 Angolan congeners) and regenerated thickened tail found in all specimens collected (n=8), a feature never recorded among Angolan congeners. It can be distinguished from the other non-Angolan western and central Africa congeners based on the same characteristics of the other Angolan species (Ceríaco et al. 2020a). Hemidactylus faustus sp. nov. can be distinguished from Angolan congeners by the thickness at the base tail. Additionally, it could be distinguished from H. mabouia by the presence of smaller subcaudal scales, and from H. benguellensis-group by lower number of precloacal-femoral pores (17–19 vs. 23–33 in H. benguellensis, and 26-28 H. cinganji sp. nov.), less keeled tubercle rows, smaller maximum length (45.3 mm [mean=39.8] vs. 54.5 [mean=47.5]), the dark dorsal uniform color with dorsolateral light stripes and absence of dorsolateral orange tubercle rows (present in H. benguellensis-group). It differs from H. carivoensis sp. nov. by the absence of keeled subcaudal scales, less keeled dorsal tubercle rows, and dark dorsal uniform color with dorsolateral light stripes and absence of dorsolateral orange tubercle rows. Hemidactylus faustus sp. nov. differs from the H. longicephalus-group by having smaller SVL (maximum length 45.3 mm [mean=39.8] vs. 60.08 [mean=46.57] in H. longicephalus and 64.8 [mean=58.96] in H. paivae), larger number of precloacal-femoral pores (17–19 vs. 6–11 in H. longicephalus and 6–8 in H. paivae) and lower number of granular scales between the dorsal tubercles (2–3 vs. 3–6 in H. longicephalus and 4–9 in H. paivae). It differs from the H. bayonii-group by having a larger SVL (maximum SVL 45.3 mm [mean=39.8]), than H. bayonii 36.2 mm [mean=34.9] and H. vernayi (42.5 mm [mean=32.7]), and lower than H. nzingae (51.5 mm [mean=44.3]) and larger number of precloacal-femoral pores (17–19 vs. 4–9 in H. bayonii, 4–6 in H. vernayi, 7–8 in H. nzingae and 8 in H. gramineus).

Measurements and meristic characters of the holotype are presented in Table S8. Adult female with a snout-vent-length (SVL) of 39.39 mm and regenerated tail length (TL) of 22.84 mm. Robust body, nape slightly distinct. Head slightly wider than the body and shorten (HW/HL 0.67). Canthus rostralis not prominent, but slightly marked. Eye diameter (3.05 mm), with vertical pupil and crenulated margin. Supraciliar scales small and pointed. Ear height (0.98 mm). Ear to eye distance slightly larger than orbit diameter (3.17 mm). Snout rounded. Frontal scales granular and of similar size as occipital scales. Occipital scales granular interspersed with large number of smooth and conical tubercle scales from eyes to nape. Rostral wider than deep (1.66 vs. 0.91 mm, respectively). Rostral semi divided posterodorsally, in contact with 1^st supralabial, nostril, two postnasal and one internasal scales. 9 supralabial and 7 infralabials. First supralabial in contact with the nostril. Nostril circular rounded by rostral, supranasal, prenasal and 2 postnasals. Prenasal, postnasal and supranasal same size. One row of scales between supralabials and the orbit. Mental large, triangular, with two larges rectangular postmental scales in short contact posteriorly to the mental. 7 post-postmental scales, composed by 2 post-postmental half size of postmental scales in contact with postmentals and 1^st and 2^nd infralabials, and 5 small post-postmentals in contact with postmental scales. Gular scales half size than ventral scales and granular. Between the gular scales and infralabials a row of enlarged scales is present, decreasing in size towards the 5^th infralabial where they become the same size as the gular scales.

Body robust and slightly short (TRL/SVL 0.41). Ventral scales widely larger than dorsal scales, with 31 scales across the belly. The dorsal pholidosis presents heterogenous conical, granular scales interspersed by 16 conical dorsal tubercle rows at midbody. Dorsal tubercle rows are separated by 3 granular scales. Tubercle scales reach the posterior part of the eye region where they lose the keeling progressively. Tubercle in the base of tail is well developed. Tail with lateral conical tubercle rows. Regenerated tail with regular larger scales (after precloacal) enlarged and 2 well-developed postcloacal spurs on each side.

Fore- and hindlimbs relatively short, stout; forearm medium sized (FL/SVL 0.23); tibia short (CL/SVL 0.18). Digits short and clawed. All digits of manus and pes indistinctly webbed. Scansors beneath toes and fingers are equally divided and composed by 1^st scansor undivided and variable number of undivided terminal scansors. Number of scansors: 6-8-7-7-7 (right manus), 7-10-11-10-9 (right pes). Relative length of digits: V < IV < III > II > I (right manus); V < IV < III > II > I (right pes).

Variation in scalation and body measurements of the paratypes and additional material of H. faustus sp. nov. are reported in Table S8. All the material examined is in concordance with the description of the holotype.

In life (specimen MNCN 50534; Fig. 16A): this species displays a nocturnal greyish or brownish coloration above with two darker dorsolateral bands from the occiput to the tail, which includes 5 W-shaped darker crossbands in contact with both lateral bands; each dark crossbar is separated by lighter blotches; head with an irregular dark and light brownish patch and a dark brownish band from the narine to anterior portion of the forelimb; light beige ventrally and laterally, with scattered black speckles in the venter; upper and lower labials beige; limbs with irregular dark-and-light brownish patches; tail with similar color and slightly banded; iris silvered with a black narrow pupil and brownish-golden reticulation. During the day, this species presents a uniform pattern along the body. In preservative (Holotype; Fig. 15): dorsum with dark coloration and five spaced darker W-shaped crossbars from the occiput to the tail that could be difficult to distinguish, with lighter dorsolateral bands; ventrum light beige with scattered black speckles. Variation: from light brownish to totally dark dorsal coloration; cross-bands can be difficult to distinguish; ventrum can be uniform beige or have scattered black speckles.

Figure 16. 

A – Dorsolateral view of Hemidactylus faustus sp. nov. (MNCN 50534). B – Pungo Andongo, habitat of H. faustus sp. nov. C – records of H. faustus sp. nov. (star depicting type locality and only known locality). Photos by Javier Lobón–Rovira.

The name “faustus” applies to a Latin word that designate ‘good luck’, evoking the serendipitous discovery of this species. The species epithet is used as a masculine adjective singular. The first specimen was found by Beatriz Vaz Pinto, daughter of PVP, under a small stone which was removed while preparing a campsite. This unexpected find led to further collecting of this new and previously unrecorded form, albeit from a locality that had been regularly surveyed since the mid-19^th century.

This species is likely a micro-endemic form, strictly associated with the conglomerate inselbergs of Pungo Andongo also known as Pedras Negras (Black Rocks), just north of the mid-Cuanza River in western Malanje Province, Angola (Fig. 16C). At the moment the only know population occurs on this site, which covers approximately 4,000 ha of huge rocky conglomerate boulders. A similar and nearby inselberg system – Pedras Jingas, albeit smaller, shares identical geological features as Pungo Andongo, being situated some 20kms to the northeast. It is quite possible that the species is present at Pedras Jingas and also in a few smaller isolated inselbergs nearby, but these areas have not been surveyed yet. All considered, it is likely that H. faustus sp. nov. is contained within about 6,000 ha of suitable habitat in the region. However, due to the limited material confirmed to belong to this species, we cannot calculate the EOO and we regard the conservation status of this species as Data Deficient. This species needs further studies about the real extent of its range and current population trends to better address its conservation status. Due to its small distribution range and highly specialized niche this species may provisionally warrant a threat status.

Hemidactylus faustus sp. nov. represents a ground-dwelling rupicolous species apparently associated with the unique geological formation of conglomerate massifs in northern Angola. It was found sheltering under small rocks or between the cavities created by plant roots growing on the flattened top of massive inselbergs, at around 1250 m a.s.l. Most specimens were collected at night foraging on the ground or at times climbing the sparse and stunted vegetation present, possibly hunting small spiders and other invertebrates. The specimens displayed an elusive behavior, jumping and disappearing quickly between the cavities and among vegetation roots when disturbed. The species occurs in sympatry with H. paivae (see Table S2 for H. paivae recorded localities), which occupies a different ecological niche, the latter living on the vertical and inaccessible walls of the conglomerate boulders. Both species could also be found in the rocky conglomerate base that makes the transition between various inselbergs. The site where the species was discovered lies within the Angolan Miombo Woodlands, even though the local ecological conditions can be considered atypical.

Hemidactylus mabouia is the most genetically distant species from all the Angolan congeners (minimum ND2 uncorrected p-distance >26%, see Table 1). Furthermore, H. mabouia can be easily differentiated from all its Angolan congeners by the presence of enlarged medial subcaudals, being absent in all Angolan Hemidactylus (Ceríaco et al. 2020a). Therefore, all records genetically or morphologically assigned to H. mabouia have been considered as unequivocal records, in absence of more accurate and recently studies within H. mabouia-group.

Consequently, this work provides 25 new records (Table S2, Fig. 17) for H. mabouia within Angolan territory. It is remarkable that this work provides the first records of H. mabouia in Bié Province, collected by Werner Conradie, Luke Verburgt and Alexander Rebelo between 2016 and 2018, and the eastern-most record within Angola territory from Lucapa, Lunda Norte Province, collected by William R. Branch and Werner Conradie in 2011 (Fig. 17). Moreover, this work demonstrates the large adaptability of H. mabouia, being present in at least 8 of the 13 AMBUs defined in this work (Fig. 17).

Figure 17. 

Records of Hemidactylus mabouia in Angola. Dark blue circles represent records provided by Ceríaco et al. 2020a; light blue circles depict new records provided by this work.